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Habitat-Dependent Transmission of Male Advertisement Calls in Bladder Grasshoppers (Orthoptera; Pneumoridae) Vanessa C

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Habitat-Dependent Transmission of Male Advertisement Calls in Bladder Grasshoppers (Orthoptera; Pneumoridae) Vanessa C The Journal of Experimental Biology 207, 2777-2786 2777 Published by The Company of Biologists 2004 doi:10.1242/jeb.01092 Habitat-dependent transmission of male advertisement calls in bladder grasshoppers (Orthoptera; Pneumoridae) Vanessa C. K. Couldridge* and Moira J. van Staaden Department of Biological Sciences, and JP Scott Center for Neuroscience, Mind and Behavior, Bowling Green State University, Bowling Green, OH 43403, USA *Author for correspondence (e-mail: [email protected]) Accepted 18 May 2004 Summary It has been hypothesized that the physical properties of fynbos species had low levels of signal attenuation over the environment exert selection pressure on long-range distance in all environments. The fynbos biome was acoustic communication signals to match the local habitat characterized by high inconsistency in signal degradation, by promoting signal characteristics that minimize excess while the forest biome had the highest levels of attenuation and distortion. We tested this in a unique environmental noise. Innate habitat characteristics, family of bladder grasshoppers notable for producing a leading to comparatively limited acoustic communication signal with a 2·km maximum transmission distance. In distances in the forest and fynbos relative to the savanna direct performance comparisons, male advertisement calls and succulent karoo, may therefore explain the need for a of seven species were broadcast through four vegetation higher quality of signal transmission in grasshoppers biomes – forest, fynbos, savanna and succulent karoo. The inhabiting the former two environments. calls of species native to forest and fynbos biomes propagated with lower levels of distortion over distance in their respective habitats relative to those of non-native Key words: acoustic communication, advertisement call, habitat, species, while fynbos species also performed best in the bladder grasshopper, Pneumoridae, sound transmission, remaining two habitats. In addition, both forest and spectrographic cross correlation. Introduction Acoustic signals are commonly utilized for communication et al., 1986), and external properties of the local environment, over long distances by a variety of animal taxa, from butterflies incorporating vegetation type (Lang, 2000), meteorological (Yack et al., 2000) to elephants (McComb et al., 2003). conditions (Garstang et al., 1995), and acoustic interference However, one of the major predicaments plaguing long-range from other organisms (Wollerman, 1999). signallers is that sounds become progressively more degraded The acoustic adaptation hypothesis emanated largely from the further they propagate. The transmission of acoustic signals the work of Morton (1975) and Hansen (1979) and was through any given environment results in modification of the conceived as a means of means of investigating the relationship signal amplitude and fidelity over distance. Loss of amplitude, between animal vocalizations and ecological factors. This resulting from spherical spreading and also from absorption hypothesis was based on the observation that environmental and scattering of sound waves (excess attenuation), diminishes factors may heavily influence the evolution of long-range the range at which receivers can detect a signal, while loss acoustic signals by imposing selection pressures that act to of fidelity, incorporating frequency-dependent attenuation, modify the sound properties of signals in order to maximize irregular amplitude fluctuations and reverberations, reduces the their broadcast range and the number of potential receivers ability of receivers to accurately recognize a signal in the event (Endler, 1992; Forrest, 1994; Morton, 1975). This could lead it is detected. A combination of changes in both amplitude and to a matching between signal and environment, such that fidelity contributes to overall patterns of signal degradation. signals transmit optimally in native habitats and, conversely, The rate of acoustic signal degradation, and subsequently perform poorly when broadcast through non-native habitats. the range at which sounds can be used effectively for Studies concerning the propagation of long-range acoustic communication purposes, is dependent upon numerous factors. signals in relation to the environment in which they are These include spectral and temporal properties of the signal transmitted have yielded mixed results. The acoustic itself (Wiley and Richards, 1978), behavioural modifications adaptation hypothesis for enhanced transmission efficiency in of the animal during signalling, such as calling from native habitats has garnered support primarily from studies preferential heights and locations (Bennet-Clark, 1987; Keuper on avian and primate taxa. Birds inhabiting different 2778 V. C. K. Couldridge and M. J. van Staaden environments produce vocalizations with characteristics suited discernable differences in the female signal and the shorter to enhancing transmission in their native habitats (e.g. Bertelli transmission distances thereof. and Tubaro, 2002; Bowman, 1979; Cosens and Falls, 1984; Pneumorids are endemic to the coastal regions of southern Gish and Morton, 1981; Handford, 1981; Morton, 1975; Africa, where they are found in four distinct vegetation biomes: Wiley, 1991). Moreover, calls broadcast in different forest, savanna, fynbos (‘fine bush’) and succulent karoo environments exhibit differential rates of excess attenuation (Rutherford, 1997). These four habitat types represent a wide and/or distortion in both birds (e.g. Dabelsteen et al., 1993; Shy range of environmental conditions, varying in extremes from and Morton, 1986) and primates (e.g. Brown et al., 1995; dense, humid, forested areas to open, arid, semi-desert areas. Waser and Brown, 1986; Waser and Waser, 1977). In contrast, The forest biome is confined mostly to the wetter eastern the influence of the environment on the evolution of signal seaboard, where it is extremely patchy in distribution and characteristics has been more difficult to establish in anurans. comprises a very small total land area. Forest vegetation at the Most studies have failed to find a significant correlation site of transmission experiments comprised a closed canopy of between habitat acoustics and either signal characteristics woody vegetation up to 25·m tall and was dominated by (Zimmerman, 1983) or signal propagation (Castellano et al., Leucosidea sericea and Ficus thonningi, with trunk diameters 2003; Kime et al., 2000; Penna and Solís, 1998; but see Ryan reaching 1·m, and an understory of Maytenus acuminata, et al., 1990). Variability in detecting the effects of habitat Hypericum and Helichrysum spp. The subtropical savanna selection are likely to be due to the multiplicity of additional biome, by far the largest biome in southern Africa, consists of factors that affect the evolution of signals. Mismatches grasses interspersed with taller woody vegetation. The area between signal and environment, which would potentially where experiments were conducted fell under the class of mask the effects of habitat selection, could arise from predator broad-leaved savanna. This is a typical C4 grassland with avoidance, competition avoidance, sexual selection, Themeda, Tristachya, Trachypogon and Aristida spp. being the phylogenetic history, and variation in the relative strengths of most prominent components, and average grass height reaching competing selection pressures. Here we test the acoustic approximately 0.5·m. The fynbos biome occupies the most adaptation hypothesis for bladder grasshoppers (Orthoptera, southern and south-western regions of southern Africa and is Pneumoridae) through an examination of acoustic signal characterized by evergreen, fine-leaved shrubs of intermediate transmission of species-specific male mating calls in different height. Fynbos vegetation is renowned for its unparalleled habitat types. This is the first detailed examination of the species diversity and is characterized by the presence of three relationship between the environment and sound propagation distinctive elements viz. restioids and ericoid shrubs, both in an invertebrate group specialized for long-distance ranging in height from 0.5·m to 2·m, and overstory proteoid signalling. We focus exclusively on habitat effects, since shrubs reaching a maximum height of 5·m (Cowling et al., constraints imposed by acoustic characteristics of the habitat 1997). The sparsely vegetated succulent karoo biome derives provide the framework within which other selection pressures its name from the low-lying leaf-succulent plants that dominate must operate. We thus anticipate an effect that is detectable, this biome. It is located along the western coast of southern though perhaps not strong. Africa and represents an intermediate state between the less arid Bladder grasshoppers comprise a small family of up to 17 fynbos to the south and true desert to the north. Succulent karoo species (Dirsh, 1965), with behaviour and morphology vegetation, although similar to fynbos in some respects, has a uniquely suited for pair-formation via acoustic duetting over relatively higher proportion of bulbous plants, and the distances of at least 50·m (van Staaden and Römer, 1997). With vegetation is lower growing, sparser, and comprises more the aid of an air-filled, inflated abdomen functioning as an succulents than fynbos. acoustic resonator, males produce a nocturnal advertisement Our primary objective was to evaluate the relative call that is both extremely loud (98·dB·SPL at 1·m) and importance
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